Arctoa (1996) 6: 113-117 NOTES ON PHILONOTIS (BARTRAMIACEAE, MUSCI). 1. STATUS AND DISTRIBUTION OF PHILONOTIS FALCATA (HOOK.) MITT. ÇÀÌÅÒÊÈ Î PHILONOTIS (BARTRAMIACEAE, MUSCI). 1. ÑÒÀÒÓÑ È ÐÀÑÏÐÎÑÒÐÀÍÅÍÈÅ PHILONOTIS FALCATA (HOOK.) MITT. TIMO KOPONEN1 ÒÈÌÎ ÊÎÏÎÍÅÍ1 Abstract Philonotis falcata (Hook.) Mitt. is exclusively an Asiatic taxon; records from North America and Europe are based on misunderstandings. It was reported from Europe in the 1820’s, but later European bryologists excluded it from the European flora. The epithet falcata was connected with Philonotis fontana (Hedw.) Brid. at the varietal level, as P. fontana var. falcata (Mitt.) Brid., and later synonymized with both P. calcarea (Bruch & Schimp.) Schimp. and P. fontana var. fontana. Philonotis falcata differs from them in many characters, such as the shape of the leaves, leaf-cell areolation, distribution of papillae and mamillae on leaf cells, and seriate leaves, and it has been placed in a different subsection. Philonotis falcata is for the first time recorded for Kirgizia, and the former Soviet Union. Ðåçþìå Philonotis falcata (Hook.) Mitt. ÿâëÿåòñÿ ÷èñòî àçèàòñêèì âèäîì. Óêàçàíèÿ íà åãî íàõîäêè â Ñåâåðíîé Àìåðèêå è Åâðîïå áûëè îñíîâàíû íà åãî íåâåðíîì ïîíèìàíèè. Åãî óêàçûâàëè â Åâðîïå â 1820-õ ãã., íî âïîñëåäñòâèè îí áûë èñêëþ÷åí èç ôëîðû Åâðîïû. Ýïèòåò falcata èìåë îòíîøåíèå è ê Philonotis fontana (Hedw.) Brid., ãäå ôèãóðèðîâàë â íàçâàíèè ðàçíîâèäíîñòè, P. fontana var. falcata (Mitt.) Brid., êîòîðóþ îòíîñèëè â ñèíîíèìû èëè ê P. calcarea (Bruch & Schimp.) Schimp., èëè ê P. fontana var. fontana. Philonotis falcata îòëè÷àåòñÿ îò ýòèõ âèäîâ ïî ìíîãèì ïðèçíàêàì, â ÷àñòíîñòè ïî ôîðìå ëèñòüåâ, ñòðîåíèå êëåòî÷íîé ñåòè, ðàñïîëîæåíèþ ïàïèëë è ìàìèëë íà êëåòêàõ ëèñòüåâ, ðàñïîëîæåíèþ ëèñòüåâ â ïðàâèëüíûõ ðÿäàõ, â ñâÿçè ñ ÷åì ñïðàâåäëèâî åãî îòíåñåíèå ê äðóãîé ïîäñåêöèè. Philonotis falcata âïåðâûå îòìå÷àåòñÿ äëÿ Êèðãèçèè è òåððèòîðèè áûâøåãî Ñîâåòñêîãî Ñîþçà. INTRODUCTION Philonotis falcata (W. J. Hook.) Mitt. For several reasons it is necessary to discuss (Figs. 1-2) the status of Philonotis falcata (Hook.) Mitt.: J. Linn. Soc. Bot. Suppl. 1: 62. 1859. - Bartra- (1) records persistently appear claiming its pres- mia falcata W. J. Hooker, Trans. Linn. Soc. Lon- ence in Europe; (2) the taxon was recently syn- don 9: 317. 27 f. 4. 1808. - P. fontana var. falca- onymized with P. calcarea (Bruch & Schimp.) ta (W. J. Hook.) Brid., Bryol. Univ. 2: 21. 1827. Schimp., which is an error, and (3) P. falcata - Type: Nepal, leg. F. Buchanan (BM!). specimens collected in Kirgizia by V. F. Broth- Hooker (1808) described Bartramia falca- erus in 1896 were located; these should have ta based on a Himalayan specimen. The de- been the first collections known from the area scription and the figures of the protologue are of the former Soviet Union, but the synonymiz- so clear that a misunderstanding is not possi- ing mentioned above postulated a wide range ble. Bridel-Brideri (1827) combined the epi- for P. falcata in European and Siberian Rus- thet to the varietal level, as Philonotis fontana sia and in Middle Asia, including Kirgizia. How- var. falcata and gave its distribution: “In tota ever, several workers have clearly shown that Europa vulgaris, uda tenens”. This obviously P. falcata is solely an East and Southeast Asi- caused long-lasting confusion. Still “Index atic taxon. muscorum” (Wijk et al. 1967) gave its distri- 1 – Department of Ecology and Systematics, P.O. Box 7, FIN-00014 University of Helsinki, Finland 114 TIMO KOPONEN bution as Europe, North America, Southeast Asia the Himalayas, China and the Philippines to and Oceania. However, several authors (see be- Japan. Kabiersch (1937) monographed East low) have already shown that P. falcata does Asiatic Philonotis and reached the same con- not belong to the European and North Ameri- clusion, as did Ochi (1962, 1963). Accordingly, can floras, and the modern checklists (Corley et it is rather well established in the literature al. 1981, Anderson et al. 1990) and floras from that P. falcata (Hook.) Mitt. (P. fontana var. these areas have excluded the taxon. falcata (Hook.) Brid.) occurs only in East and Düll (1985) proposed that P. fontana var. Southeast Asia, reaching Hawaii (Bartram 1933). falcata (Hook.) Brid. is a synonym of P. calcar- Ignatov & Afonina (1992) followed Düll’s ea (Bruch & Schimp.) Schimp., and Anderson (1985) synonymization of P. falcata with P. et al. (1990) synonymized it with P. fontana calcarea, and since they accepted the latter tax- var. fontana, giving no reason, however, for the on at the varietal level, their P. fontana var. synonymizations. Warnstorf (1906) discussed the falcata (Hook.) Brid. gained a wide distribu- similarity and difficulties in separating falcate- tion in arctic and western Europe, Caucasus, leaved forms of P. fontana from sterile P. cal- Siberia, and Central Asia. carea: “Var. falcata Schpr., Synops. ed. II, p. 520 TAXONOMY (1876) [= Schimper 1876] ist eine kräftige, an Ochi (1962) explained very clearly why Ph. calcarea erinnernde Form ..” “Eine ähnli- Philonotis falcata can never be confused with P. che Form, wie sie in Kryptogamenfl. v. Deutschl. fontana and its relatives; they have different types IV. Bd. 2. Abt. p. 569 [= Limpricht 1895] als var. of distribution of leaf cell mammillae and papil- falcata Brid. beschrieben wird.... und es dürfte lae. The papillae in P. falcata are exclusively sit- vergebliche Mühe sein, dieselbe von steriler Ph. uated at the distal end of the leaf cell, but in P. calcarea unterscheiden zu wollen.” fontana the mamillae or papillae are at the proxi- Loeske (1906) supported Warnstorf’s view: mal end of the leaf cells, or rarely in the middle of “Warnstorf hat in seinem neuen Mooswerke zum- the cells if the cells are very short. Other charac- ersten Male eine var. falcata so beschrieben, dass ters separating P. falcata and P. fontana are the sie nur als form der Ph. fontana gedeuted wer- leaf shape, and the size of cells in different parts den kann. Diese Form muss Ph. fontana var. fal- of leaves. P. falcata has very distinctly carinate cata Br. eur. ex p. Warnst. oder einfach var. fal- leaves with costa strongly bending, and the leaves cata Warnstorf heissen”. By this statement Loeske are broadest at the insertion and gradually taper- (1906) actually created a later homonym P. ing to an acute or acuminate apex. The leaf cells fontana var. falcata Warnst. ex Loeske., which are long and narrow down near the base, where may have increased the confusion. Warnstorf’s several rows of cells a little wider and shorter (1906) treatment was accepted by Dismier are present; short acute leaves may, however, have (1908); he cited falcate-leaved forms of P. fon- short and wide cells throughout the leaf. Such tana “var. falcata (Br. eur. et Syn. musc., ex p.) plants Ochi (1962, 1963) included in the variety Warnst., Krypt., II, p. 611, 1905”. It is obvious carinata (Mitt.) Ochi. P. fontana has a wide ovate that these European authors, M. G. Dismier, K. G. leaf base from which the apical part is rather Limricht, L. Loeske, W. P. Schimper, and C. Warn- abruptly narrowed to an acute or acuminate apex. storf, excluded the Asiatic type of P. fontana P. fontana has a distinct area of wide cells in the var. falcata (Hook.) Brid. from their concept of lower part of the leaf, and the cells in the nar- P. fontana var. falcata, and Kabiersch (1937) rowed apical part are much narrower than these followed them. lower cells. Dismier (1910) later studied American One characteristic usually present in Philono- Philonotis and did not even mention P. falca- tis falcata and absent from P. fontana is the ta in his paper. Moreover, he (Dismier 1912) seriate position of the leaves. In P. falcata the made a study on P. falcata and its related leaves are in distinct rows, best seen on leaves on taxa in which he synonymized many names and innovations, while P. fontana has no rows. Ka- gave the distribution in detail. The western-most biersch (1937) emphasized this difference be- specimen seen by him came from Afghanistan, tween P. falcata and P. fontana by placing wherefrom the taxon extends its range through them in different subsections, Falcatae Kab. and Status and distribution of Philonotis falcata (Hook.) Mitt. 115 1 mm B A D C E 100 μm F G 3 mm Fig. 1. Philonotis falcata (Hook.) Mitt. (from Brotherus reliquiae 10231, H): a – plant; b – leaves; c – leaf border near apex; d – leaf cells at mid-leaf; e – leaf border at midleaf; f – basal leaf cells; g – angular leaf cells. Scale bars: 3 mm – for a; 1 mm – for b; 100 μm – for c-g. Fontanae Kab. (nom. illeg., = subsection Philono- the leaves not being basal on cells (except in tis). The characteristics of subsection Falcatae P. seriata). I think that the subsection Falcat- are leaves mostly in rows (in P. seriata Mitt., ae, if limited as Kabiersch (1937) proposed, is P. falcata and in P. turneriana (Schwaegr.) not natural. On the basis of their leaf cell mor- Mitt. in Kabiersch’s sense) and the papillae of phology and the distribution of their papillae, 116 TIMO KOPONEN Fig. 2. Distribution of Philonotis falcata (Hook.) Mitt. Specimens in H and H-BR. P. seriata and P. yezoana Card. & Besch. ex in it may represent another taxon with a wide Card. are more closely related to P. fontana distribution in the Pacific and Melanesia. than to P. falcata. Philonotis calcarea, with which Figure 2 is a preliminary map of the distri- P. falcata was synonymized by Düll (1985) and bution of Philonotis falcata. Worth reporting Ignatov & Afonina (1992), is closely related to are the first collections of P. falcata from the P. fontana. Kabiersch (1937) discussed P.